Biological Control
Compost Tea as a Container Medium Drench
for Suppressing Seedling Damping-Off Caused by Pythium ultimum
Steven J. Scheuerell and Walter F. Mahaffee
First author: Department of Botany and Plant Pathology, Oregon State University; and second author: U.S. Department of Agriculture–
Agriculture Research Service-Horticulture Crops Research Laboratory, Corvallis, OR 97330.
Accepted for publication 14 June 2004.
ABSTRACT
Scheuerell, S. J., and Mahaffee, W. F. 2004. Compost tea as a container
medium drench for suppressing seedling damping-off caused by Pythium
ultimum. Phytopathology 94:1156-1163.
Compost tea is being used increasingly in agricultural production to
control plant diseases. However, there has been limited investigation re-
lating disease control efficacy to various compost tea production methods,
particularly compost tea produced with active aeration and additives to
increase microbial population densities in compost tea. Aerated compost
tea (ACT) and nonaerated compost tea (NCT), produced with or without
additives, was investigated for the suppression of damping-off of cucum-
ber caused by Pythium ultimum. Compost tea was used to drench soilless
container medium inoculated with P. ultimum; effect on damping-off
Compost tea is being used increasingly as an alternative plant
disease control measure in commercial horticulture (13). Compost
tea is produced by mixing compost with water and incubating for
a defined period, either actively aerating (aerated compost tea,
ACT) or not (nonaerated compost tea, NCT) and with or without
additives that are intended to increase microbial population
densities during production (13,14). Compost tea applied to
foliage has been demonstrated to suppress a range of foliar dis-
eases (reviewed in literature citations 14 and 19); however, the
use of compost tea as a soil drench for seed or root rot suppres-
sion has received very little attention.
The control of damping-off disease of seedlings, commonly
caused by Pythium spp. in Northern latitudes, is of particular
interest to greenhouse growers (15,16). Damping-off can be a
severe problem when peat-based medium, which is naturally con-
ducive to the pathogen, is used (5). The only investigation to date
involving Pythium spp. and compost tea determined that pea
seeds soaked in NCT, dried, and sown 2 days later had reduced
disease symptoms on seedlings caused by P. ultimum (17). Heat
treating the NCT negated all suppression of pathogen growth in
vitro, indicating the likely role of the NCT microflora in disease
suppression (17).
The microflora of both NCT (19) and ACT (6) are typically de-
scribed as being dominated by bacteria, and therefore the bac-
terial population of compost tea could be a useful parameter to
measure in relation to plant disease suppression. It has been
proposed that increasing the population of total and active bac-
teria in ACT will generally increase the level of plant disease sup-
Corresponding author: W. F. Mahaffee
E-mail address: mahaffew@science.oregonstate.edu
Publication no. P-2004-0901-01R
This article is in the public domain and not copyrightable. It may be freely re-
printed with customary crediting of the source. The American Phytopathological
Society, 2004.
1156 PHYTOPATHOLOGY
ranged from not suppressive to consistently suppressive depending on the
method used to produce the tea. The most consistent formulation for
damping-off suppression was ACT produced with kelp and humic acid
additives. Producing ACT with a molasses-based additive inconsistently
suppressed damping-off; evidence suggests that residual nutrients can in-
terfere with disease suppression. Heating or diluting compost tea negated
suppression. Across all compost tea samples, there was no significant
relationship of bacterial populations, measured as active cells, total cells,
or CFU, to disease suppression. However, for all ACT produced without
the molasses-based additive, there was a threshold of bacterial population
density (6 log10 active cells per ml, 7.48 log10 total cells per ml, or 7 log10
CFU per ml) above which compost teas were suppressive.
pression (6). However, there is no published evidence supporting
these assertions.
The objectives of this study were to determine if compost tea,
produced from commercially available composts, applied as a
drench suppresses seedling damping-off of cucumber caused by
P. ultimum in a peat-based container medium that is naturally
conducive to the disease; to examine how producing compost tea
with aeration and additives impacts disease suppression; and to
determine if the level of disease suppression is related to bacterial
population density (measured as active cells, total cells, and
colony forming units [CFU]) in compost teas.
MATERIALS AND METHODS
Producing compost teas. Three commercial sources of com-
post were used to make compost teas (Table 1). Yard trimmings
compost (Rexius, Inc., Eugene, OR) was produced from ground
landscape trimmings in windrows turned weekly for 3 months and
then cured in a large pile for 9 months before sampling. One
cubic yard of this material was cured an additional 2 years before
being used for these experiments. Vermicompost was produced
from mixed vegetation in a vertical flow reactor and marketed in
10-kg polyethylene bags (Soil Soup, Inc., Edmonds, WA). After
receiving the bagged product, the compost was cured for 2 months
in a shaded, open container. Tea compost is a proprietary blend of
vegetative and animal manure-based composts sold for making
compost tea (Rexius, Inc.). Approximately one cubic yard of the
tea compost was cured for 5 months in a shaded, open container.
A Bio-blender (Soil Soup, Inc.) was used to produce ACT;
mechanical aeration and liquid circulation was done by air in-
jection through a submersed, hollow propeller shaft. Fifteen liters
of tap water at 20 to 22°C was placed in a 19-liter bucket and
aerated for 2 h to reduce chlorine present in the water. If used,
additives were added to the water (Table 2). Compost inoculum
was added to the liquid by immersing compost (500 g, approxi-
mately 50% moisture, wt/wt) held in a 100-µm mesh filter bag
(Soil Soup, Inc.). To assist the removal of soluble material and
microorganisms from the compost, the filter bag was lifted above
the water, allowed to drain into the bucket for 15 s, and then re-
immersed for 30 s. This was performed a total of three times, with
the filter bag left suspended in the liquid. The Bio-blender ran
continuously for the remainder of the 36-h production cycle.
The NCT was started by adding 15 liters of tap water at 20 to
22°C to a 19-liter bucket, and the water was allowed to stand for
24 h for passive chlorine removal. If used, additives (Table 2)
were added to the water followed by pouring 500 g of compost
into the water. The entire contents were stirred vigorously for 20 s
and then left undisturbed for 7 to 9 days until used.
Chemical and biological properties of compost and compost
teas. Chemical properties. The pH and electrical conductivity
(EC) were recorded twice for the vermicompost and tea compost
and four times for the yard trimmings compost during the course
of the study. Compost pH was determined from a saturated paste
with a portable pH meter (Model 150; IQ Scientific Instruments,
San Diego, CA) and EC was determined from a 2:1 (vol/vol) mix-
ture of distilled water and compost with a portable EC meter
(Model 933100; Hanna Instruments, Woonsocket, RI) (9). For
compost teas, the pH, EC, temperature, and dissolved oxygen
(Model 600; Engineered Systems & Design, Newark, DE) were
recorded for each batch by immersing the probes into the bucket
just before use in the P. ultimum bioassay.
Microbiological populations of compost. A 10-g sample of
compost was added to 90 ml of sterile 0.02 M potassium phos-
phate buffer (PPB), pH 7.0, in a 250-ml shaker flask, and shaken
(300 rpm, 25°C) for 20 min. The samples were serially diluted,
plated using an automated spiral plater (Eddy Jet; IUL Instru-
ments, Barcelona, Spain) onto selective agar, and incubated at
22°C. Bacteria were enumerated on 5% trypticase soy broth agar
(1.5 g of Difco trypticase soy broth and 15 g of agar per liter with
100 µg/ml cycloheximide [TSBAcyc100]). Fungi were enumerated
on water agar, pH 6 (18 g of agar per liter with 50 µg/ml rifampi-
cin [WArif50]). Yeast was enumerated on dilute, selective yeast
media (SYM; 1.5 g of yeast extract, 2.5 g of peptone, 5 g of dex-
trose, 2.3 g of malt agar, and 17 g of agar per liter) amended with
100 µg/ml chloramphenicol, 50 µg/ml ampicillin, 500 µg/ml
streptomycin sulfate, and 2 µg/ml dichloran. Population densities
are reported as CFU/g of dry compost.
Microbiological populations of compost teas. A 1-ml sample of
compost tea was aseptically removed from each bucket at the end
of the culture period. Following serial dilution in sterile PPB,
dilutions were plated on TSBAcyc100, WArif50, and SYM, and
then incubated and counted as described previously. Populations
were recorded as CFU/ml of compost tea.
Active and total bacterial cells in compost tea. Active and total
bacterial cells were enumerated by epifluorescent microscopy and
sequential digital imaging using the following procedure. Deter-
mination of metabolically active cells was done by staining with
fluorescein diacetate (FDA; Sigma-Aldrich, St. Louis, MO), and
determination of total cells was done with 4,6-diamidino-2-
phenylindole (DAPI; Sigma-Aldrich). Stock solutions of DAPI
(0.2 mg/ml DAPI in sterile deionized H2O) and FDA (0.2 mg/ml
TABLE 2. Additives used to make compost tea
Additive recipe Additive components per liter of water
No additive None
Bacterial 5 ml of Bacterial Nutrient Solution (Soil Soup Inc.,
Edmonds, WA)
Fungalz 1.2 g of Maxicrop soluble seaweed powder (Maxicrop
USA Inc., Arlington Heights, IL)
2.5 ml of Humax liquid humic acids (JH Biotech Inc.,
Ventura, CA)
3 g of rock dust (Target Glacial Dust; Target Products
Ltd., Burnaby, B.C., Canada)
z
Adapted from Ingham and Alms (7).

TABLE 1. Chemical and biological properties of compost and compost teas

Population density (log10)s
Compost Compost Dissolved Filamentous Bacterial population
tea tea oxygenr Tempera- ECr Yeastt fungiu
Compostn aerationo additivesp N
q
(ppm) turer (°C) pHr (ds/m) (CFU) (CFU) CFUv Active cellsw Total cellsx
Yard trimmings 4 ... ... 5.7 ± 0.1y 0.87 ± 0.08 5.10 ± 0.11 5.25 ± 0.14 7.96 ± 0.21 ... ...
NCT None 5 6.4 ± 0.5 15.5 ± 2.0 6.7 ± 0.3 0.26 ± 0.05 bdlz 0.69 ± 1.4 5.27 ± 0.26 5.31 ± 0.59 6.45 ± 0.42
NCT Bacterial 5 0.2 ± 0.1 15.2 ± 1.9 5.3 ± 1.0 0.95 ± 0.13 2.17 ± 2.5 3.84 ± 0.10 6.18 ± 0.51 5.76 ± 0.49 7.49 ± 0.32
NCT Fungal 5 0.2 ± 0.1 15.3 ± 2.3 6.9 ± 1.1 1.05 ± 0.08 0.73 ± 1.5 1.39 ± 1.6 7.14 ± 0.38 6.11 ± 0.39 7.26 ± 0.45
ACT None 6 8.5 ± 0.5 18.1 ± 1.6 7.4 ± 0.2 0.27 ± 0.03 bdl 0.70 ± 1.2 6.11 ± 0.41 5.84 ± 0.34 6.96 ± 0.40
ACT Bacterial 9 8.0 ± 0.4 18.5 ± 2.0 7.8 ± 0.5 0.96 ± 0.18 2.37 ± 1.6 0.58 ± 1.1 8.20 ± 0.38 6.98 ± 0.33 8.18 ± 0.36
ACT Fungal 13 8.2 ± 0.4 18.6 ± 2.6 8.6 ± 0.3 1.00 ± 0.11 bdl 1.05 ± 1.3 7.82 ± 0.22 6.87 ± 0.34 7.78 ± 0.47
Vermicompost 2 ... ... 6.0 ± 0.8 4.70 ± 0.2 4.70 ± 0.02 6.09 ± 0.16 8.33 ± 0.33 ... ...
ACT None 3 9.0 ± 0.6 17.8 ± 1.1 7.3 ± 0.5 0.77 ± 0.02 bdl 1.85 ± 1.6 6.02 ± 0.30 5.74 ± 0.06 7.20 ± 0.51
ACT Bacterial 4 8.6 ± 0.7 18.1 ± 1.1 8.0 ± 0.9 1.51 ± 0.23 2.86 ± 1.9 bdl 8.74 ± 0.16 7.35 ± 0.37 8.42 ± 0.28
ACT Fungal 4 8.7 ± 0.7 17.8 ± 1.9 8.5 ± 0.1 1.62 ± 0.18 1.50 ± 1.7 1.23 ± 1.4 7.84 ± 0.19 7.03 ± 0.25 8.11 ± 0.50
Tea compost 2 ... ... 6.9 ± 0.2 3.52 ± 0.02 4.60 ± 0.64 6.32 ± 0.20 9.13 ± 0.16 ... ...
ACT Bacterial 2 8.3 ± 0.6 17.3 ± 2.5 8.0 ± 0.4 1.93 ± 0.10 3.53 ± 0.21 2.10 ± 3.0 8.60 ± 0.28 6.92 ± 0.02 8.17 ± 0.24
ACT Fungal 2 8.2 ± 0.9 16.1 ± 1.0 8.5 ± 0.0 1.67 ± 0.02 bdl 1.97 ± 2.8 8.04 ± 0.49 6.86 ± 0.480 8.61 ± 0.50
n Yard trimmings compost (Rexius, Inc., Eugene, OR); vermicompost, vegetative-based vermicompost sold for compost tea (Soil Soup, Inc., Edmonds, WA); and
tea compost, proprietary compost blend sold for compost tea use (Rexius, Inc.).
o ACT = aerated compost teas; NCT = nonaerated compost tea.
p Additives–none, bacterial, fungal, and bact-fungal fermentation additives described in Table 2.
q Number of batches.
r Recorded at end of ACT (36 h) and NCT (7 to 10 days) fermentation period.
s CFU/dry g for compost; CFU/ml for compost tea yeast, fungi, and CFU of bacteria; cells/ml for compost tea active cells and total cells.
t Colony forming units of yeast enumerated on dilute, selective yeast media (1.5 g of yeast extract, 2.5 g of peptone, 5 g of dextrose, 2.3 g of malt agar, and 17 g of

agar per liter), amended, per milliliter, with 100 µg of chloramphenicol, 50 µg of ampicillin, 500 µg of streptomycin sulfate, and 2 µg of dichloran.
u Water agar (pH 6.0) with 100 ppm of rifampicin (CFU/dry g for compost; and CFU/ml for compost tea).
v Trypticase soy broth agar (5%) with 100 ppm of cycloheximide (CFU/dry g for compost; and CFU/ml for compost tea).
w Fluorescein diacetate stained after membrane filtration, cells per milliliter of compost tea.
x 4,6-Diamidino-2-phenylindole (DAPI) stained after membrane filtration, cells per milliliter of compost tea.
y Mean ± standard deviation.
z All samples below detection limit of log 2.3 CFU/ml of compost tea.
10

Vol. 94, No. 11, 2004 1157

FDA in dimethyl sulfoxide) were kept frozen at –20°C with fresh
stock used each day. A working solution of FDA was made by
adding 1 ml of stock solution to 9 ml of PPB. For staining, an
appropriate 10-ml suspension of compost tea was prepared from
the serial dilutions used for plating on media. The DAPI stock
solution (100 µl) was added to 10 ml of compost tea resulting in
0.002 mg of DAPI per ml. After 3 min incubation in darkness, the
sample was vacuum-filtered through a black 0.2-µm filter (Isopore
Membrane Filter 0.2-µm GTBP02500, 25-mm diameter, Milli-
pore, Bedford, MA), on top of a support pad (MF Support Pad,
AP1002500, 25-mm diameter, Millipore) in a 25-mm diameter
glass microanalysis filter (#0-97563G, Fisher Scientific, Pittsburg,
PA), which was covered with foil to reduce light exposure
throughout the process. After filtration, the vacuum was stopped.
Sterile PPB (1 ml) was gently overlaid on the filter, allowed to sit
for 5 s, and vacuum filtered through the filter. The FDA working
solution (1 ml) was overlaid on the filter, allowed to sit for 2 min,
and then vacuum-filtered through the filter. The filter was
immediately adhered to a glass slide with a 20-µl drop of sterile
PPB and examined microscopically.
Digital imaging of stained cells was done at ×400 (Lieca
DMRB Microscope, Heerbrugg, Switzerland) with 480/40-nm ex-
citation and 510-nm LP barrier filters (GFP cube) and 360/40-nm
excitation and 420-nm barrier filters (UV cube) to observe FDA-
and DAPI-stained cells, respectively. The filter was viewed and a
digital image was captured (Spot RT Slider diagnostic camera
with Spot version 3.1 image capture software 2000 [Diagnostic
Instruments, Inc., Sterling Heights, MI]) using the GFP cube, a
second image was captured using the UV-cube. The stage was
moved arbitrarily and the process was repeated until six pairs of
images were obtained. Image acquisition was completed before
visible quenching of the FDA-stained cells occurred.
Each pair of images were merged with the UV-cube image used
as the source image and GFP-cube image as the active image, and
semi-automated counting of the total number of stained cells was
performed using Image Pro Plus version 4.1 software (Media
Cybernetics, Silver Springs, MD). The counted cells were verified
and manually recounted if necessary. The active cells, FDA
stained (green), were manually counted from the merged image.
The process was repeated for each pair of images and the num-
ber of active and total cells were averaged across the six image
pairs. To determine the number of cells per milliliter of compost
tea, the average number of active and total cells from the six
images was multiplied by 3,341.995973 to extrapolate the number
of stained cells in the camera frame area to the exposed filter area
(2.1 cm2) and multiplied by the dilution factor.
Damping-off suppression by compost-amended container
media or compost tea drench. Compost-amended container
media. The three compost sources (described previously) were
tested individually for their ability to suppress P. ultimum damp-
ing-off of cucumber when incorporated into container medium
that was inoculated with P. ultimum and drenched with water.
Three compost-amended container media were prepared by mix-
ing compost (1:3, vol/vol) with commercial peat-perlite growing
medium (Sunshine Mix #1, Sun Gro Horticulture, Inc., Van-
couver, B.C., Canada).
Growing medium for compost tea. Compost tea drench treat-
ments were tested for the ability to suppress P. ultimum damping-
off of cucumber in 100% commercial peat-perlite growing
medium (Sunshine Mix #1, Sun Gro Horticulture, Inc.) that was
inoculated with P. ultimum.
Cucumber seedling assay. P. ultimum (isolated from corn roots,
Willamette Valley, OR; provided by B. Hoinacki) inoculum was
produced using 14-day cultures growing on Ko and Hora’s (8)
soil and chopped potato medium, dried, and sieved through a
1-mm2 grid. Particles retained on a 0.25-mm2 sieve were used to
inoculate growing medium. For each compost-amended container
medium or compost tea drench treatment, 2 liters of the respective
1158 PHYTOPATHOLOGY
growing medium was thoroughly mixed with P. ultimum inocu-
lum (1.0 g/liter of growing medium) in a Twin Shell Dry Blender
(Patterson-Kelley Co., East Straudsberg, PA) for 2 min. Inocu-
lated medium was placed evenly into six 400-ml plastic nursery
pots (experimental units). Eight cucumber seeds (Cucumus
sativus cv. Marketmore 76) were sown 1 cm deep into each of the
pots. For each compost tea drench treatment, compost tea was
applied to the six replicate pots using a fine-spray watering can
until the pots were saturated. Compost-amended container media
were similarly moistened with tap water. The six pots for each
treatment were placed, in random order, on separate nursery trays
that served as experimental blocks. A large, clear, plastic bag was
inflated and sealed around each nursery tray to simulate a
germination room and maintain even moisture in the pots. The
flats were placed in a 20°C growth chamber with a 16-h photo-
period. At 3 and 6 days after planting (DAP), each flat was vented
to minimize changes in the atmosphere within the sealed flats. At
9 DAP, the number of healthy seedlings was recorded for each
pot. A seedling was classified as healthy if it was growing nor-
mally and had no symptoms or signs of infection. Infection symp-
toms included a water-soaked or yellowing stem, wilted cotyle-
dons, and stem lesion leading to seedling collapse; pathogen sign
was white mycelia covering any portion of the seedling.
Effect of heating and diluting compost tea on disease devel-
opment. The cucumber seedling assay was used to test whether
heating or diluting compost tea impacted Pythium suppression.
Compost tea was heated with continuous stirring to 95 to 98°C
for 30 min and cooled to 25°C before applying to seeded pots as
previously described. Diluted compost tea was prepared by mix-
ing compost tea 1:1, 1:4, and 1:9 (vol/vol) with tap water before
applying to seeded pots as previously described.
Effect of residual nutrients on disease development. The cu-
cumber seedling assay was used to test whether the presence of
excess nutrients from additives used to produce compost tea im-
pacted Pythium suppression. Nutrient concentration in compost
teas was manipulated in three separate ways. First, ACT was
produced with 0.5, 1.0, and 1.5% bacterial additive (Table 2) with
the assumption that not all of the available nutrients in the higher
nutrient concentration treatments would have been used during
the 36-h production period. Second, ACTs made with or without
the fungal additive were amended with 0.1% (vol/vol) bacterial
additive just prior to being used as a drench. Third, 0.01, 0.04,
0.125, and 0.3% molasses was added (vol/vol) to ACT made with
the fungal additive immediately before drenching. The last two
experiments simulated excess nutrients at quantified levels.
Experimental design and statistical analysis. All experiments
were designed as randomized complete blocks with nursery flats
as blocks and individual pots as the experimental unit. Each ex-
periment had both pathogen-inoculated and noninoculated peat-
perlite growing medium control treatments that were drenched
with tap water. All experiments were repeated at least once.
Two-way analysis of variance was performed with treatment
and block as factors; means were separated from the water drench
control using mean separation tests such as Fisher’s protected
least significant difference, Duncan’s multiple range, or paired t
tests as indicated and appropriate. Linear regression was used to
relate compost tea bacterial populations to the percent healthy
cucumber seedlings. Percent healthy seedlings for each treatment
were calculated by dividing the treatment mean healthy seedlings
by the mean healthy seedlings of the noninoculated peat-perlite
control treatment in each seedling assay. All statistical analyses
were done with Statgraphics 4.0 software (Manugistics, Rock-
ville, MD).
RESULTS
Chemical and biological properties of compost and compost
teas. The pH, EC, and culturable populations of bacteria, yeast,
and filamentous fungi varied across the three compost sources
(Table 1). In compost tea, the level of dissolved oxygen was influ-
enced by the use of aeration and additives during production
(Table 1). The average dissolved oxygen content across all ACT
was 8.4 ppm. For compost tea made from the yard trimmings
compost, the average dissolved oxygen content of NCT without
additives was 6.4 ppm, significantly lower than 8.5 ppm for ACT
made without additives (P < 0.0001). Adding either the bacterial
or fungal additives to NCT decreased dissolved oxygen to 0.2 ppm.
These NCT with greatly reduced oxygen conditions released
highly putrescent odors upon drenching. Compost tea temperature
was not influenced by production method; for all preparations,
compost tea temperatures closely followed ambient temperatures
(data not shown). Compost tea pH varied across the different
compost tea production methods (Table 1). When ACT samples
across compost sources were grouped by additive type, the
average pH of compost tea made with no nutrient (pH 7.4), with
bacterial additive (pH 7.9), and with fungal additive (pH 8.6)
were significantly different from each other (P < 0.0001). The EC
of all ACT produced without additives (0.40 ds/m) was signifi-
cantly lower (P < 0.0001) than both ACT made with the bacterial
additive (1.23 ds/m) and with the fungal additive (1.02 ds/m).
Microbial communities of both NCT and ACT were predomi-
nantly bacteria (Table 1), with most bacteria occurring as indi-
vidual planktonic cells (data not shown). The average population
of culturable bacteria, active bacterial cells, and total bacterial
cells increased with the use of additives during compost tea pro-
duction across compost sources (Table 1). While fungal popula-
tions were significantly lower than the source compost in all com-
post tea preparations, the highest culturable population densities
of fungi were recovered from NCT produced with the bacterial
additive (Table 1). A surface mat of organic material and microbial
biomass partially consisting of sporulating filamentous fungi
formed in these open, static buckets. However, there were no sig-
nificant differences (P = 0.46) in the mean population of cultur-
able fungi between the bacterial-additive and fungal-additive
amended compost teas across all ACT samples (Table 1). In both
NCT and ACT made without additives, the average culturable
populations of yeast (CFU/ml) were at least 250-fold (vermicom-
post) and 629-fold (yard trimmings compost) lower than the
source compost (CFU/dry g), while the average culturable fungi
(CFU/ml) were 17,379-fold (vermicompost) to 36,291-fold (yard
trimmings compost) lower than the source compost (CFU/dry g)
(Table 1). Adding the bacterial additive consistently increased the
average yeast population above levels found in compost teas
made without additives (Table 1).
TABLE 3. Comparison of methods to estimate bacterial populations in compost teass made with or without aeration and additives
Population density of bacteria (log10/ml)
Aerationt Additive componentsu Nv CFUw
ACT None 15 6.3 ± 0.42z
ACT Bacterial 20 8.5 ± 0.4
ACT Fungal (kelp, humic acids, rock dust) 24 7.9 ± 0.1
ACT Rock dust or humic acids 4 6.0 ± 0.29
ACT Kelp or kelp and humic acids 4 7.7 ± 0.15
NCT None 10 5.8 ± 0.71
NCT Bacterial 10 6.7 ± 0.47
NCT Fungal (kelp, humic acids, rock dust) 10 7.2 ± 0.4
s All compost teas were made with yard trimmings compost, vermicompost, or tea compost; not all samples were used in the Pythium ultimum assays presented in
this study.
t ACT = aerated compost tea; NCT = nonaerated compost tea.
u Fungal and bacterial additive components described in Table 2.
v Sample size.
w CFU/ml of compost tea; medium, 5% trypticase soy broth agar amended with 100 ppm of cycloheximide.
x Cells/ml; fluorescein diacetate stained after membrane filtration.
y Cells/ml; 4,6-diamidino-2-phenylindole (DAPI) stained after membrane filtration.
z Mean ± standard deviation.
The bacterial population of compost tea measured as colony
forming units, active cells, or total cells was influenced by the use
of aeration and additives during production. Estimations of cultur-
able populations were generally statistically equal or significantly
greater (P < 0.05) than populations measured as active or total
cells when additives were added to ACT (Table 3). For both ACT
and NCT made without additives, the average bacterial culturable
population was equivalent (P > 0.1) to the active cell population,
while the total cell population was significantly greater (P <
0.001) than the culturable population (Table 3).
However, NCT made with either the bacterial or fungal addi-
tives had significantly (P < 0.001) greater total populations and
significantly (P < 0.01) lower active populations compared with
that of culturable populations (Table 3). With active aeration and
addition of fungal additive, the total cell population was not dif-
ferent (P > 0.1) from the culturable population. With active aera-
tion and bacterial additive, the culturable population was signifi-
cantly greater (P < 0.05) than the total cell population (Table 3).
Based on data for ACT produced with components of the fungal
additive, adding kelp with or without the humic acids resulted in a
similar relationship of bacterial colony forming units to active and
total cells compared with the complete fungal additive recipe
(kelp, humic acids, and rock dust) (Table 3).
Damping-off suppression by compost-amended container
media or compost tea drench. When the yard trimmings com-
post, tea compost, or vermicompost was individually mixed (1:3,
vol/vol) with the peat-perlite growing medium, damping-off
caused by P. ultimum was significantly (P < 0.05) suppressed in
one bioassay by the yard trimmings compost (Table 4). In con-
trast, these conducive composts were individually used to produce
compost teas, that when used as a drench resulted in damping-off
that ranged from not suppressive to consistently suppressive
depending on the method used to produce compost tea. Severe
damping-off was observed when NCT containing yard trimmings
compost and no additive was used as a drench; disease severity
was not significantly (P > 0.05) different from the inoculated con-
trol in five repeated bioassays (data not shown). Drenching with
NCT produced with yard trimmings compost and either bacterial
additive or fungal additive significantly (P < 0.05) suppressed
damping-off in four of five and three of five repeated bioassays,
respectively (data not shown).
Producing ACT without additives or with the bacterial additive
resulted in inconsistent damping-off suppression over repeated
bioassays (Table 4). When ACT was produced with a combination
of bacterial and fungal additives, damping-off was not suppressed
over three bioassays (data not shown). Producing ACT with any
Paired t test P value
Bacterial CFU Bacterial CFU
Active cellsx Total cellsy vs. active cells vs. total cells
6.1 ± 0.37 7.2 ± 0.17 0.12 0.0002
7.3 ± 0.23 8.3 ± 0.09 0.0001 0.005
7.0 ± 0.16 8.0 ± 0.24 0.0001 0.23
5.7 ± 0.16 6.9 ± 0.19 0.16 0.02
6.0 ± 0.3 7.5 ± 0.36 0.01 0.41
5.8 ± 0.87 6.7 ± 0.57 0.69 0.0004
6.0 ± 0.19 7.8 ± 0.51 0.01 0.0001
6.4 ± 0.17 7.6 ± 0.26 0.0007 0.001
Vol. 94, No. 11, 2004 1159
compost and fungal additive alone suppressed damping-off in 19
of 19 P. ultimum bioassays (Table 4). When mixed with water and
drenched, the additive formulations did not suppress damping-off
(data not shown). Suppression of damping-off was not limited to
peat-perlite growing medium. Suppression was observed with
ACT produced with fungal additive in composted fir bark/peat-
perlite growing medium (1:1, vol/vol) that was inoculated with
P. ultimum (data not shown).
The bacterial population metrics of ACT were positively related
to each other; linear regression of colony forming units and total
cells (P < 0.0001; R2 = 0.67) and linear regression of colony form-
ing units and active cells (P < 0.0001; R2 = 0.77) were significant.
The relationship between percentage of healthy seedlings and bac-
terial population metrics of compost tea greatly depended on the
types of compost tea that were considered. When all ACT were
pooled, there was no linear relationship between the bacterial popu-
lation metrics and plant health; linear regression analyses of the
percent healthy seedlings of ACT against bacterial colony forming
units, active cells, and total cells explained only 13.2, 7.07, and
7.77% of the variation, respectively. Likewise, when only ACT
made with the bacterial additive was considered, based on
regression analyses, there was no relationship (P > 0.1) between
percentage of healthy seedlings and the bacterial population.
However, when ACT made with the bacterial additive were
excluded from the analysis, a significant (P < 0.05) positive linear
relationship was observed between the bacterial populations and
plant health (Fig. 1).
For the 18 NCT samples tested in bioassays (data not shown),
no positive linear relationship existed between the average num-
ber of healthy seedlings and any bacterial population metric.
Linear regression of healthy seedlings to bacterial colony forming
units, total cells, and active cells had an R2 of 0.13, 0.08, and
0.00001, respectively.
Effect of heating and diluting compost tea on disease
development. Heating NCT or ACT to 95 to 98°C for 30 min and
cooling to 25°C before drenching significantly (P < 0.05) in-
creased damping-off compared with that of unheated compost tea
(Table 5). Diluting compost tea 1:9 (vol/vol) with tap water sig-
nificantly (P < 0.05) reduced disease suppression in five of six
trials, with intermediate dilution rates having variable reductions
in suppression (Table 5).
Effect of residual nutrients on disease development. When
producing ACT, increasing the bacterial additive concentration
from 0.5% to 1.0 or 1.5% (vol/vol) significantly (P < 0.05) de-
creased the number of healthy cucumber seedlings (Fig. 2),
whereas mixing 0.1% (vol/vol) bacterial additive with suppressive
ACT just before drenching negated disease suppression (Fig. 3).
Since molasses constituted one-third of the bacterial additive by
volume, the role of unused molasses in enhancing disease devel-
opment was investigated. Mixing from 0.01 to 0.3% molasses
with suppressive ACT just before drenching significantly (P <
0.05) reduced suppression (Fig. 4). In the absence of P. ultimum,
the healthy seedlings resulting from drenching with either bac-
terial additive (0.5%) or molasses (0.3%) in water or compost tea
was not different from drenching with 100% water (data not
shown).
DISCUSSION
Drench application of compost tea can suppress P. ultimum
damping-off of cucumber in soilless container media. Initial re-
search on the use of compost teas for plant disease suppression
utilized NCT that was generally produced without additives (19);
however, recent attention has advocated the use of aeration and
additives (6). In this study, both ACT and NCT significantly re-
duced disease; but consistent disease reduction was only attained
with ACT produced with specific additives (i.e., kelp and humic
acid). While NCT produced with yard trimmings compost and
additives can suppress P. ultimum damping-off of cucumber,
relatively long production times, and putrescent odors associated
with NCT made with additives, detract from their utility. In ad-
dition, inconsistent disease control observed with NCT produced
with yard trimmings compost and additives was observed also in
preliminary bioassays using NCT produced with other compost
sources (data not shown).
The most consistent formulation for damping-off suppression
was ACT fermented with kelp, humic acids, and rock dust
(termed fungal additive). The suppressiveness of this formulation
was significantly reduced by heat treatment or dilution with tap
water. Further characterization indicated that the rock dust com-
ponent was not necessary for damping-off suppression and caused
excessive wear on mechanical parts (data not shown). Suppres-
sion conferred with the fungal additive was independent of the
compost source used in ACT production. This indicates that the
selection of additives was more critical than the source of com-
post for producing ACT that suppressed P. ultimum damping-off.

TABLE 4. Capacity of compost-amended container media or aerated compost tea drench treatments to consistently suppress Pythium ultimum damping-off of
cucumber seedlings in repeated bioassays
Yard trimmings compost Tea compost Vermicompost
Proportion of Proportion of Proportion of
Compost amendments or Compost tea suppressive Healthy suppressive Healthy suppressive Healthy
Compost tea drencht additiveu bioassaysv seedlingsw bioassaysv seedlingsw bioassaysv seedlingsw
Compost incorporated 25% by volumes 1/3 2.44 ± 2.27 0/2 0.91 ± 0.35 0/3 2.50 ± 0.44
ACTx None 1/6 2.31 ± 0.51 ndy nd 1/3 3.28 ± 0.48
ACT Bacterial 3/9 3.14 ± 1.53 0/2 2.92 ± 0.59 3/4 3.27 ± 1.99
ACT Fungal 13/13 4.85 ± 0.77 2/2 5.25 ± 1.06 4/4 4.76 ± 0.97
P. ultimum-inoculated controlz 0/13 1.43 ± 0.57 0/2 0.83 ± 0.71 0/4 1.46 ± 0.55
Uninoculated control 13/13 6.96 ± 0.53 2/2 6.75 ± 0.12 4/4 6.71 ± 0.34
s Yard trimmings compost (Rexius Inc., Eugene, OR); vermicompost = vegetative-based vermicompost sold for compost tea (Soil Soup, Inc., Edmonds, WA); and
tea compost = proprietary compost blend sold for compost tea use (Rexius, Inc.). Compost mixed with peat-perlite growing medium (1:3, vol/vol; inoculated
with P. ultimum and water drenched).
t Drenches applied to 100% peat-perlite growing medium inoculated with P. ultimum.
u Listed in Table 2.
v Proportion of significantly suppressive bioassays of total bioassays (significantly greater mean number of healthy seedlings than P. ultimum-inoculated control,

according to least significant difference test, P = 0.05; determined separately for each bioassay).
w Mean number of healthy seedlings averaged over bioassays ± standard deviation. Treatments were applied to six replicate pots with eight seeds each sown in P.

ultimum-inoculated peat-perlite growing medium. Mean number of healthy seedlings was determined by summing the number of healthy seedlings for each pot
(0 to 8) and averaging these six values.
x ACT = aerated compost tea.
y Not determined.
z Peat-perlite growing medium inoculated with P. ultimum and drenched with water.

1160 PHYTOPATHOLOGY
This has important practical implications for widespread pro- drenching with water. In addition, all growing medium had a pH
duction of disease-suppressive compost tea because additives of 6.3 to 6.5 at the end of the 9-day cucumber bioassay (data not
are standardized materials, whereas the properties of composted shown). Based on laboratory analysis of the well-cured yard
materials can vary depending on fluctuations in feedstock trimmings compost, 13 ppm of NH4 was present. Therefore, less
materials, microbial decomposition dynamics, and environmental than 1 ppm of NH4 from the compost would be in solution after
conditions. diluting approximately 25-fold (vol/vol) with water in compost
Across all compost tea samples produced with or without tea production. Lastly, ACT made with a combination of the fun-
additives, there was no significant relationship of bacterial popu- gal and bacterial additives had a pH of 8.4 and did not suppress
lations, measured as active cells, total cells, and CFU, to disease damping-off (data not shown). In total, these data support the
suppression. However, when compost teas produced with the ad- conclusion that disease suppression afforded by compost tea
dition of molasses were removed, there was a threshold of bac- applications is related to microflora present in the tea.
terial population level above which compost teas were suppres-
sive. The transition from nonsuppressive compost tea drenches to
suppressive drenches occurred at approximately 6 log10 active
bacterial cells per milliliter. The transition to suppressive samples
as measured by total bacterial cells per milliliter is less dramatic.
However, when samples with average healthy seedlings above
50% of the noninfested control were arbitrarily considered to be
suppressive, then 15 of 16 samples with greater than 7.5 log10
total cells per milliliter would be considered suppressive. Using
the same arbitrary definition of suppression, all compost teas that
had greater than 7 log10 CFU per ml of bacteria would be con-
sidered suppressive. Therefore, it appears that ACT made without
molasses-based additive can be divided into suppressive or nonsup-
pressive based upon threshold of bacterial populations as meas-
ured by active cells, total cells, and CFU (e.g., 7 log10 CFU/ml).
Based on these data, monitoring bacterial populations could be a
useful indicator for generating Pythium-suppressive compost teas
and determining if suppressive compost tea could be diluted
before use and still remain effective.
The use of aeration and additives in compost tea production
appears to determine the proportion of the total bacterial popu-
lation that was culturable. For NCT produced with or without
additives, the population of total bacterial cells was significantly
greater than the population determined by dilution plating, indi-
cating the majority of cells would not grow under the culture
conditions imposed. The same was true for ACT made without
additives or when ACT made with rock dust or humic acids were
considered as a group. Aggregation of bacterial cells could be a
cause for lower culturable populations; however, extensive aggre-
gation was not evident when observing samples for total and ac-
tive cell enumeration (data not shown). For the above types of
compost tea, the bacterial population, based on dilution plating,
underestimates the total bacterial cell population and the domi-
nant bacterial types might not be readily culturable.
However, in ACT made with the bacterial or fungal additives,
bacterial populations as colony forming units were statistically
equivalent or greater than the total bacterial cells. Having greater
culturable populations than total populations is difficult to con-
ceive. It is likely due to the observation that a slightly greater
density of bacteria was localized at the outer edge of the stained
filters (data not shown) and the image acquisition procedure did
not account for this nonuniformity. Despite this phenomenon, it
appears that either ACT made with the bacterial or fungal addi-
tives selectively increased culturable bacteria. These findings in-
dicate that the simpler culturing procedure can be used to monitor Fig. 1. Relationship of bacterial populations of aerated compost teas (ACT) to
the total bacterial population in ACT produced with readily healthy cucumber seedlings in Pythium ultimum damping-off bioassays. For
available additives, and that it is possible to readily culture the each compost tea drench treatment, healthy seedlings were scaled to per-
centage of noninoculated control (percent healthy seedlings = treatment mean
dominant bacterial strains from these compost teas.
healthy seedlings/noninoculated control mean healthy seedlings). The ACT
Since the ACT fermented with fungal additive had a pH of 8.5, with bacterial additive was not included. All ACT was produced either without
the damping-off suppression observed in this study could have additives or with single components, binary combinations of components, or
been due to reduced saprophytic activity of P. ultimum by in- all fungal additive components (0.12% [wt/vol] powdered soluble kelp, 0.25%
creasing the pH of the growing medium. Increasing soil pH to 8.5 [vol/vol] humic acids, and 0.30% [wt/vol] glacial rock dust). There was a
can cause increasing release of NH4, which is known to suppress significant linear relationship (P < 0.05 with R2 = 0.36, 0.22, and 0.57 for A,
B, and C, respectively. A, Active bacterial cells were measured by staining
Pythium spp. activity (12). However, it is unlikely that disease
with fluorescein diacetate. B, Total bacterial cells were measured by staining
suppression is primarily due to pH, since drenching with either with 4,6-diamidino-2-phenylindole (DAPI). C, Bacterial colony forming units
the fungal additive in water (pH 8.6) or heat-treated fungal were cultured on 5% trypticase soy broth agar amended with 100 ppm of
additive ACT (pH 8.5) resulted in the same disease incidence as cycloheximide.

Vol. 94, No. 11, 2004 1161

TABLE 5. Effect of diluting and heating compost tea on seedling health
Dilution ratiow
Compost tea
Compostt aerationu Additivesv 1:0x 1:1 1:4 1:9 Heat-treatedy
Yard trimmings ACT None 36.8 bz ... ... ... 15.8 a
Yard trimmings ACT Bacterial 51.2 b ... ... ... 0.0 a
Yard trimmings ACT Fungal 73.6 b ... ... ... 7.9 a
Yard trimmings ACT Fungal 61.5 b ... ... ... 20.5 a
Yard trimmings ACT Fungal 86.0 b ... ... ... 25.6 a
Yard trimmings ACT Fungal 63.6 b 34.1 a 31.8 a 25.0 a ...
Yard trimmings ACT Fungal 96.9 b 91.1 b 70.3 ab 44.2 a ...
Vermicompost ACT Fungal 74.3 cd 52.3 a–c 40.4 a–c 33.3 ab 19.0 a
Yard trimmings NCT Bacterial 70.4 b ... ... 15.1 a ...
Yard trimmings NCT Bacterial 85.3 b ... ... 10.4 a 18.3 a
Yard trimmings NCT Bacterial 86.9 b ... ... ... 39.5 a
Yard trimmings NCT Fungal 57.8 b ... ... 49.9 b 13.1 a
t Compost source described in Table 1.
u ACT = aerated compost tea; NCT = nonaerated compost tea.
v Additives described in Table 2.
w Compost tea was diluted with tap water at a given ratio.
x Compost tea was used without dilution or heat treatment directly from production bucket.
y Compost tea was heated to 95 to 98°C for 30 min and then cooled to 25°C for drenching.
z Healthy seedlings scaled to percentage of noninoculated control (% healthy seedlings = treatment mean healthy seedlings/noninoculated control mean healthy

seedlings). Numbers within rows followed by the same letter are not significantly different (P = 0.05, Duncan’s multiple range test).

Fig. 2. Effect of molasses-based, bacterial additive concentration (Soil Soup,

Inc., Edmonds, WA) used to make aerated compost tea on seedling health and
bacterial populations. Active and total cells determined by direct counting
after staining with fluorescein diacetate and DAPI, respectively. Populations
as colony forming units were determined on 5% trypticase soy broth agar with
100 ppm of cycloheximide. Compost tea drench treatments were applied to six
replicate pots each with eight cucumber seeds sown in Pythium ultimum-
infested peat-perlite growing medium. Mean number of healthy seedlings was
determined by summing the number of healthy seedlings for each pot (0 to 8)
and averaging these six values. Letters labeling bars indicate significantly
different number of healthy seedlings, and treatments with the same letter are
not significantly different (P = 0.05, Duncan’s multiple range test).
Fig. 3. Effect of tank mixing molasses-based Bacterial Nutrient Solution (Soil
Soup, Inc., Edmonds, WA) with aerated compost tea (ACT) on seedling health.
Compost tea drench treatments were applied to six replicate pots each with
eight cucumber seeds sown in Pythium ultimum-infested peat-perlite growing
medium. Mean number of healthy seedlings was determined by summing the
number of healthy seedlings for each pot (0 to 8) and averaging these six values.
Fungal ACT drench was produced with 0.12% (wt/vol) soluble kelp, 0.25%
(vol/vol) humic acids, and 0.30% (wt/vol) glacial rock dust. Bars with the same
letters are not significantly different (P = 0.05, Duncan’s multiple range test).

There are several possible explanations why repeated experi-
ments using ACT made with the molasses-based bacterial additive
had erratic damping-off suppression. The most probable explana-
tion is that residual nutrients, most likely sucrose, varied across
compost tea batches and this enhanced Pythium propagule germi-
nation and growth or reduced competition for seed exudates, re-
sulting in enhanced pathogen growth and infection. It is generally
accepted that bacteria can indirectly protect seeds against Pythium
infection by metabolizing seed-exudate stimulants (18) and that
competition for available nutrients is a likely mechanism of
soilborne pathogen suppression if adding nutrient amendments
negates suppression (10). Evidence supporting this conclusion
includes the inverse relationship between the concentration of
bacterial additive used in ACT production and disease suppres-
sion; ACT produced with increasing concentrations of sucrose as
the sole fermentation nutrient corresponded to increased damp-
ing-off disease levels (data not shown); and the addition of as
little as 0.01% molasses or 0.1% bacterial additive to suppressive
ACT significantly increased damping-off. Similarly, suppression
of cucumber seedling damping-off caused by P. ultimum in com-
post-amended container medium was negated by adding 0.375%
sucrose and 0.075% asparagine to the growing medium (3).
Residual nutrients could also suppress antibiotic production or
parasitic activity in favor of saprophytic metabolism. This would
affect bacteria capable of producing antimicrobial compounds
that reduce Pythium germination through fungistatic or fungicidal
effects (18). Excessive nutrients have been shown to reduce hyphal
lysis of P. aphanidermatum by antagonistic bacteria in separated

1162 PHYTOPATHOLOGY

Fig. 4. Effect of tank mixing molasses with aerated compost tea (ACT) on
seedling health. Compost tea drench treatments were applied to six replicate
pots each with eight cucumber seeds sown in Pythium ultimum-infested peat-
perlite growing medium. Mean number of healthy seedlings was determined
by summing the number of healthy seedlings for each pot (0 to 8) and
averaging these six values. The fungal ACT drench was produced with 0.12%
(wt/vol) soluble kelp, 0.25% (vol/vol) humic acids, and 0.30% (wt/vol) glacial
rock dust. Bars with the same letters are not significantly different (P = 0.05,
Duncan’s multiple range test).
cattle manure-compost medium (11). Amending the container
medium with a glucose-asparagine mixture (3.36:1, wt/wt),
amended at 0.5% wet weight, reduced hyphal lysis to 18% over a
24-h period compared with 80% lysis for nonamended medium
(11). Additionally, drenching the glucose-asparagine mixture (1%
solution in water) onto the compost medium negated cucumber
damping-off suppression (11).
While further work is needed to directly quantify the residual
sucrose concentrations in ACT produced with molasses-based
additive to determine the effect on damping-off suppression, there
are strong indications that the use of simple sugars as additives
should be avoided when producing compost tea for disease sup-
pression. In addition to the potential of residual nutrients increas-
ing Pythium damping-off, the use of simple sugars in producing
compost tea has been linked to growth of E. coli in aerated
compost tea makers when compost contaminated with E. coli was
used (1). Other work determined that Salmonella enterica and
E. coli O157:H7 did not grow when inoculated into nonaerated
flasks containing 20 g of compost and 180 ml of sterile water;
however, incremental additions of molasses were related to in-
creasing growth of both organisms (4). Other unpublished work
suggests that using additives with compost that is free of human
pathogens does not pose a safety risk (13). However, commercial
testing of compost for human pathogens cannot guarantee a
pathogen-free product due to potential sampling errors and detec-
tion limits of common analytical procedures (13). Further work is
needed to characterize additives that sufficiently increase micro-
bial populations for plant disease control yet do not increase
human or plant pathogens (13).
An increased understanding of the quantitative relationship be-
tween compost tea microbial populations and plant disease devel-
opment could help develop guidelines for producing suppressive
drench formulations, similar to guidelines for compost-amended
container medium that prescribe minimum levels of microbial ac-
tivity and biomass to prevent P. ultimum damping-off (2). Infor-
mation on the duration of suppression afforded by drenching con-
tainer medium with compost tea would be useful to develop an
application schedule for long-term suppression of Pythium damp-
ing-off in commercial production. Further experimentation on a
production scale is needed to develop this method into an effec-
tive disease control tool.
ACKNOWLEDGMENTS
We acknowledge Soil Soup, Inc., for supplying Bio-Blenders for
producing aerated compost tea. We thank B. Short for technical assistance
and B. Hoinacki for the P. ultimum culture. Funding for this research was
provided by USDA CRIS 303-5358-22000-024-00D and the Oregon
Association of Nurseries and Oregon Department of Agriculture.
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